.

Структурно-функціональні адаптації крові коропа до екстремальних факторів водного середовища: Автореф. дис… канд. біол. наук / О.С. Смольський, НАН

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IAOe?IIAEUeIA AEAAeAI?ss IAOE Oe?a?ie

?INOEOOO A?Ae?IA?IEIA??

Niieuenueeee Ieaenaiae? Na?a?eiae/

OAeE 577.41/46 + 597.554

NO?OEOO?II-OOIEOe?IIAEUeI? AAeAIOAOe?? E?IA? EI?IIA AeI
AENO?AIAEUeIEO OAEOI??A AIAeIIAI NA?AAeIAEUA

03.00.17 – a?ae?ia?ieia?y

A A O I ? A O A ? A O

aeena?oaoe?? ia caeiaoooy iaoeiaiai nooiaiy

eaiaeeaeaoa a?ieia?/ieo iaoe

Ee?a – 1998

Aeena?oaoe??th ? ?oeiien.

?iaioa aeeiiaia a eaai?aoi??? a?io?i?? oaa?ei eaoaae?e o?i??
*a?i?a?anueeiai aea?aeaaiiai iaaeaaia?/iiai oi?aa?neoaoo ?iai? O. A.
Oaa/aiea

Iaoeiaee ea??aiee – aeieoi? a?ieia?/ieo iaoe, aeioeaio

A?oa?iei Aaneeue Aaneeueiae/,

Oa?iii?euenueeee aea?aeaaiee iaaeaaia?/iee

oi?aa?neoao ?iai? Aieiaeeie?a Aiaothea,

Ia?oee i?i?aeoi?, caa?aeoaa/
eaoaae?ith

caaaeueii? a?ieia??

Io?oe?ei? iiiiaioe – aeieoi? a?ieia?/ieo iaoe,

NIEIIAO?IA Aaeaioeia Aeieo??aia,

?inoeooo a?ae?ia?ieia?? IAI
Oe?a?ie, i?ia?aeiee iaoeiaee ni?a?ia?oiee a?aeae?eo aeieia?/ii?
o?c?ieia?? aiaeyieo oaa?ei

eaiaeeaeao a?ieia?/ieo iaoe,

Eo?aio Aieiaeeie? Caiia?eiae/,
Oa?iii?euenueeee

aea?aeaaiee iaaeaaia?/iee
oi?aa?neoao

?iai? Aieiaeeie?a Aiaothea,
aeioeaio

eaoaae?e o?i??

I?ia?aeia onoaiiaa – Ee?anueeee Iaoe?iiaeueiee oi?aa?neoao ?iai? Oa?ana
Oaa/aiea, a?ieia?/iee oaeoeueoao,

I?i?noa?noai ina?oe Oe?a?ie

Caoeno aeena?oaoe?? a?aeaoaeaoueny ” 2 ” ethoiai 1999 ?. i 1000
aiae ia can?aeaii? niaoe?ae?ciaaii? a/aii? ?aaee Ae 26.213.01. i?e
?inoeooo? a?ae?ia?ieia?? IAI Oe?a?ie ca aae?anith: 254210, i. Ee?a-210,
i?iniaeo Aa?i?a Noae?ia?aaea, 12

C aeena?oaoe??th iiaeia iciaeiieoeny a a?ae?ioaoe? ?inoeoooo
a?ae?ia?ieia?? IAI Oe?a?ie, i. Ee?a, i?iniaeo Aa?i?a Noae?ia?aaea, 12

Aaoi?aoa?ao ?ic?neaiee ” 24 ” a?oaeiy 1998 ?.

A/aiee nae?aoa?

niaoe?ae?ciaaii? a/aii? ?aaee

eaiaeeaeao a?ieia?/ieo iaoe
Ni??iiaa I.I.

CAAAEUeIA OA?AEOA?ENOEEA ?IAIOE

Aeooaeuei?noue oaie. Aaaeeeaith neeaaeiaith i?iaeaie aea/aiiy iaoai?ci?a
aio?iiiaaiii? aeaa?aaeaoe?? aiaeieo aeinenoai oa ?ic?iaee iiaeo iaoiae?a
a?ieiio?ieth ?o ??aiy ? aeeiai?ee ? aeine?aeaeaiiy ?aaeoe??
iieaeoey?ieo oa iaoaaie?/ieo nenoai i?aai?ci?a ia ae?th e?i?ooth/eo
aeieia?/ieo oaeoi??a aiaeiiai na?aaeiaeua. A?aeiii, ui iaei??th c
iae/ooeea?oeo oa oaeaeei?aaaoth/eo ia ?o ci?io oeaiei i?aai?cio ? e?ia.
Inoaii?i /anii ia aaiaoieia?/iee aiae?c, yeee aeicaiey? ca
ii?oi-ooieoe?iiaeueieie oa a?io?i?/ieie oa?aeoa?enoeeaie e?ia?
ia’?eoeaii ioe?ieoe oa ia?aaeaa/eoe iayai?noue /e ?icaeoie no?ano
caaeiaai aei i?iyao ee?i?/ieo iciae iaoieia??, caa?oa?oueny nooo?aa
oaaaa (Na?ioi?i, 1991). Iniaeeai oe?iiei ? oeae i?aeo?ae i?e ioe?ioe?
?ioieneeaoe?e (Eioia, 1991, Oae?o/aiei oa ?i., 1988). Aeine?aeaeaiiy
iieaeoey?ieo iaoai?ci?a no?oeoo?ii? aaeaioaoe?? eiiiiiaio?a e?ia? i?e
?ioieneeaoe?yo, ye cacia/a? Naiai/aaa A.I. (1972), aeicaieeoue ?ice?eoe
iaoai?cie caaacia/aiiy ooieoe?iiaeueiiai aiiainoaco e?ia?. A ca’yceo c
oeei aea/aiiy aaeaioeaieo ?aaeoe?e nenoaie e?ia? a?ae?ia?iio?a oa iiooe
oeo ?ioaa?aeueieo iieaciee?a e?ia?, ye? aeacothoue ia eeth/ia? ci?ie
noaio ?o i?aai?cio ca aeno?aiaeueiiai aieeao na?aaeiaeua, ? aeooaeueiei
aeey ioe?iee oiene/iiai caa?oaeiaiiy aiaeieo aeinenoai oa a?iiii?oi?eiao
noaio ? ?iaeoeo ?ioieneeaoe?e a?ae?ia?iio?a.

Iaoa oa caaaeaiiy aeine?aeaeaiue. Iaoith ?iaioe aoei aea/aiiy
o?c?ieiai-a?io?i?/ieo iaoai?ci?a ?aaoeyoe?? ooieoe?iiaeueii? aeoeaiino?
aaiiaeia?iiai? nenoaie oa iniaeeainoae aia?aaoe/iiai iaoaaie?cio e?ia?
?ea a oiiaao no?ano, aeeeeeaiiai ae??th aa?ioe/ieo ? aio?iiiaaiieo
oaeoi??a aiaeiiai na?aaeiaeua, oa ?ic?iaea ? aaeineiiaeaiiy ia oe?e
iniia? iaoiae?a iii?oi?eiao aiaeieo aeinenoai oa ?ioieneeaoe?e
i?aai?cio ?ea.

A caaaeaiiy aein?aeaeaiue aoiaeeei aea/aiiy neeaaeo oa
o?c?ieiai-a?io?i?/ieo oa?aeoa?enoee e?ia? ?ea a oiiaao ceiiaiai
aieiaeoaaiiy, i?e ci?i? ?I aiaeiiai na?aaeiaeua oa ca ae?? oieneeaio?a.
Aeine?aeaeoaaee ai?no oi?iaieo aeaiaio?a e?ia?; eiioeaio?aoe?th oi?i
aaiiaeia?io; no?oeoo?i? iiaeeoeeaoe?? a?ee?a e?ia? oa aaoa?iaaii?noue
aaiiaeia?io; e?iaoeeo iane/aiiy aaiiaeia?io eeniai oa aeeiai?eo
eeneioii? ? eoaeii? aeaiaoo?aoe??; ??aaiue aeoeaiino? oa?iaioieo nenoai
neioaco oa iaoaaie?cio aaio ? aia?aaoe/iiai iai?io a a?eo?ioeeoao;
ai?no a e?ia? oinoao?a, aeeth/ii iae?ia?a?/ieo, ye iieaciee?a
aia?aaoe/iiai noaio e?ia? ? ?aaoeyoi??a ooieoe?iiaeueii? aeoeaiino?
aaiiaeia?iiai? nenoaie ca ae?? oaeoi??a aiaeiiai na?aaeiaeua.

Iaoeiaa iiaecia ?iaioe. Aia?oa i?iaaaeaii eiiieaenia aeine?aeaeaiiy
?aaeoe?? e?ia? a oiiaao i?e?iaeiiai no?ano, aeeeeeaiiai ceiiaei
aieiaeoaaiiyi ca iecueeeo oaiia?aoo?, oa ca ae?? aeno?aiaeueii? ci?ie
eeneioiino? na?aaeiaeua ? oieneeaio?a i?aai?/ii? (oaiie) ? iai?aai?/ii?
(?iie a?iaaiieo oa iaa?iaaiieo iaoae?a) i?e?iaee ca ?iaeea?aeoaeueiiai
oa ni?eueiiai (a noi?oao) aieeao. Anoaiiaeaii caaaeuei? caeiiii??iino?
oi?ioaaiiy eiiiainaoi?ii-aaeaioeaii? a?aeiia?ae? aaiiaeia?iiai?
nenoaie. Aia?oa iieacaii oi?aa?naeuei?noue iaoai?cio i?aeo?eiaiiy
no?oeoo?ii-ooieoe?iiaeueii? oe?e?ni?no? aaiiaeia?io i?e no?aniaeo
aieeaao ??cii? i?e?iaee ca ?aooiie ai?aeoaaiiy oa ae?eoaaiiy eiai
iieaeoee. Aeine?aeaeaii iaoai?ci ?aaoeyoi?iiai aieeao ia
ooieoe?iiaeueiee noai aaiiaeia?io e?ia? ?ea iae?ia?a?/ieo oinoao?a oa
nenoai aia?aaoe/iiai iaoaaie?cio a?eo?ioeeo?a. Aeyaeaii iaai?
ei?aeyoe?ei? caeaaeiino? i?ae aeine?aeaeaieie ??aiaaeie iieacieeaie
e?ia?, ia iniia? /iai c?iaeaii aeniiaie i?i oa, ui ?acenoaioi?noue oa
aaeaioeai?noue ?ea aei aeno?aiaeueieo oaeoi??a aecia/a?oueny an?i
niaeo?ii iioaioe?eii ?nioth/eo eiiiainaoi?ii-aaeaioeaieo iaoai?ci?a
caoenoo ooieoe?iiaeueieo oa?aeoa?enoee e?ia?.

I?aeoe/ia oe?ii?noue ?iaioe. Ia iniia? iaea?aeaieo ?acoeueoao?a aeyaeaii
eeth/ia? iieacieee e?ia?, ye? iiaeia aeei?enoaoe aeey ioe?iee noaio oa
?icaeoeo ?ioieneeaoe?e a i?aai?ci? ?ea, a oaeiae aeey iii?oi?eiao
caa?oaeiaiue aiaeiiai na?aaeiaeua: ni?aa?aeiioaiiy e?aaiaeieo oi?i
aaiiaeia?io; ei?aeyoe?y e?iaoeee aeenioe?aoe?? ieneaaiiaeia?io c ??aiai
AOO a a?eo?ioeeoao e?ia?; ??aaiue noeueoa?ae?eeueiuo a?oi oa nooi?iue
ai?aeoaaiiy ? ae?eoaaiiy a?ee?a e?ia? (aaiiaeia?io oa a?ee?a ieacie) ye
oaeoi?a ae?aioaaiiy ai?iieeneioieo ?aaeeeae?a a?ae oieneeaio?a,
iniaeeai, ?ii?a aaaeeeo iaoae?a oa oieneeaio?a ieenithaaeueii? ae??.

Ai?iaaoe?y ?iaioe. Iniiai? iieiaeaiiy ?iaioe aeiiia?aeaeeny ia:

1. I?aeaoc?anuee?e iaoeiai-i?aeoe/i?e eiioa?aioe?? (Oe?a?ia, *a?i?a?a,
*AeI? ?i. O.A. Oaa/aiea, 1995).

2. I?aeaoc?anuee?e iaoeiai-i?aeoe/i?e eiioa?aioe?? (Oe?a?ia, Ee?a, OAAI,
1996).

3. 7 Oe?a?inueeiio a?io?i?/iiio c’?cae? (Oe?a?ia, Ee?a, 1997).

4. 2 c’?cae? A?ae?iaeieia?/iiai oiaa?enoaa Oe?a?ie (Oe?a?ia, Ee?a,
1997).

5. 1 I?aeia?iaei?e iaoeiai-i?aeoe/i?e eiioa?aioe?? “Aeieia?y e iieiaeue”
(?anioae?ea Aaea?onue, Aiiaeue, 1998).

6. 1 Anaoe?a?inuee?e iaoeia?e eiioa?aioe?? « Aeieia?/iee no?an ?
aaeaioaoe?y a a?ieia?/ieo nenoaiao» (Oe?a?ia, Oa?iii?eue, 1998).

Ioae?eaoe??. Ii iaoa??aeao aeena?oaoe?? iioae?eiaaii 13 ?ia?o.

No?oeoo?a e ia’?i ?iaioe. Aeena?oaoe?y neeaaea?oueny ?c anooio, iaeyaeo
e?oa?aoo?e (1 ?icae?e), aenia?eiaioaeueii? /anoeie (3 ?icae?ee),
i?aenoie?a ? aeniiae?a. ?iaioa aeeeaaeaia ia 168 noi??ieao, i?noeoue
25 ?enoie?a, 9 oaaeeoeue, 220 e?oa?aoo?ieo aeaea?ae, c yeeo 102 –
ia ?iicaiieo iiaao.

Aeaeea?aoe?y eiie?aoiiai aianeo. An? iaea?aeai? ?acoeueoaoe aeine?ae?a,
?o iaaiai?aiiy ? aeniiaee aeeiiai? aeena?oaioii iniaenoi.

IAEssAe E?OA?AOO?E

I?iaiae?ciaai? o?c?ieiai-a?io?i?/i? ci?ie a e?ia? ?ea ca ae??
??cieo oaeoi??a aeno?aiaeueiiai oa?aeoa?o, a oaeiae ca ??cieo oiia
?nioaaiiy. Ia iniia? oeeo aeaieo c?iaeaii aeniiaie i?i iaiao?aei?noue
eiieaeniiai aea/aiiy a?aeiieo oa anoaiiaeaiiy iiaeeeaeo ca aeaieo oiia
?nioaaiiy iaoai?ci?a no?oeoo?ii-ooieoe?eii? a?aeiia?ae? aaiiaeia?iiai?
nenoaie e?ia? ?ea oa ?ic?iaee iiaeo iaoiae?a a?ieiio?ieth oiene/ieo
noai?a a i?aai?ci? a?ae?ia?iio?a.

2. Iaoa??aee ? iaoiaee aeine?aeaeaiue

Ia’?eoii aeine?aeaeaiiy aoee ei?iiia? ?eae (Cyprinus carpio L.) ??/iiai
a?eo ianith 120-150 a, yeeo aeo?eioaaee a i?e?iaeieo noaaeao ca
ii?iaeueiiai aaciaiai oa a?ae?io?i?/iiai ?aaeei?a. Aeine?aee i?iaaaeaii
a aeaa??oiao ia’?iii 200 e, a ye? ?ici?uoaaee ii 6 iniaei ?ea. Ia??iae
aee?iaoe?? noaiiaea 14 ae?a. Ai?no eenith eieeaaany a iaaeao 5,0-7,0
ia/e. Aiaeo a aeaa??oiao ci?ithaaee ?ac a /ioe?e aeiae. Iaoe??ieo
caoaeiee?a ia?aceoa?ieo oai?ia oa no??/eiaeo ia?aceo?a a ?ea ia
aeyaeaii.

Aea/aee ii?oi-ooieoe?iiaeueiee noai e?ia? ei?iia ca ae?? oaiieo, ?ii?a
iaoae?a (iaai?th, i?ae?, ia?aaioeth oa naeioeth) ie?aii oa a noi?oao c
ai?aeii. ?iie a?ae?a?aii ca i?eioeeiii a?ieia?/ii? ae?? (a?iaaii? oa
iaa?iaaii?). Aeine?aeaeaiiy aieeao ai?aeo i?iaiaeeee a ca’yceo c oei, ui
a?i, aoaeo/e iaoaaie?oii oaa?ei, o aenieeo eiioeaio?aoe?yo aeyaey?
oiene/io ae?th. Ine?eueee iniiaiei ia’?eoii aeine?aeaeaiue ? aaiiaeia?i,
ooieoe?iiaeueia aeoeai?noue yeiai cia/ii caeaaeeoue a?ae aioo??oiuei-
oa ciai?oiueia?eo?ioeeoa?iiai ?I, aea/aii oaeiae aieea ci?ie eeneioiino?
na?aaeiaeua (?I 6,4 + 0,1 oa 8,1 + 0,1).

Oiene/i? ??ai? (2 ?eaiainiiaea?nueeeo AAeE) ?a/iaei o aiae? aeaa??oi?a
noai?thaaee aianaiiyi ?o ?ic?aooieiaeo e?eueeinoae. Oiene/io
eiioeaio?aoe?th ai?aeo a aiae? (0,1 ia/e – ii?iaiaee ??aaiue)
i?aeo?eioaaee aianaiiyi ai?aeiai? aooa?ii? noi?o? NH4OH: NH4Cl c ?I
7,2; Pb2+ (0,2 ia/e) – aoeaoaoo naeioeth; Cu2+ (0,2 ia/e) –
noeueoaoo i?ae?; Zn2+ (2,0 ia/e) – noeueoaoo oeeieo; Mg2+ (100,0 ia/e)
– oei?eaeo iaai?th c a?aooaaiiyi i?e?iaeii? oaa?aeino? aiaee; Mn2+ (2,41
a/e) – noeueoaoo ia?aaioeth; oaiieo (0,002 ia/e) – ?ic/eiaiiai
na?aeiia?aaiaiiai oaiieo. Eeneioiee oa eoaeiee no?an noai?thaaee
aianaiiyi o aiaeo HCl /e NaOH. Cia/aiiy ?I uiaeiaiai eiio?iethaaee ?
ei?aaoaaee.

E?ia a?aeae?aee ?c neiona cya?iai? aaie c aeei?enoaiiyi ye
aioeeiaaoeyioo aaia?eio c ?ic?aooieo 10 iae/ie e?ia?. E?euee?noue
oi?iaieo aeaiaio?a e?ia? aecia/aee a eaia?? Ai?y?aa (Aoieinueeee ?
?i., 1990); aaiaoie?eo – oeaio?eooaoaaiiyi a aaiaoie?eoi?e oeaio?eooc?
IOeA-8; caaaeueiee ai?no aaiiaeia?io ? eiai oi?i ?ic?aoiaoaaee ca
aeeoa?aioe?aeueieie niaeo?aie iiaeeiaiiy (E?eeeeaee ? ?i., 1979).
?ic?aooiie ni?aa?aeiioaiiy iene-, aeaciene- oa iao- oi?i aaiiaeiaeio
cae?enithaaee ca oi?ioeaie Ymai K. et al. (1970), Benesh R. E. et al.
(1973). Aeoeai?noue i?oioiiae??eii? d – ai?iieaaoeeiaoneioaoace ia/?iee
aecia/aee ca Granick S. (1966), a aeoeai?noue IAAeH – caeaaeii?
iaoaaiiaeia?i?aaeoeoace e?ia? – ca Aea?a?c A. A. (1975). E?iaoeeo
eoaeii? oa eeneioii? aeaiaoo?aoe?? aaiiaeia?io aea/aee ca Noa?iaeoa I.
O. (1979). Nii??aeiai?noue aaiiaeiaeio aei eenith ioe?ithaaee ca
e?eaeie aeenioe?aoe?? ieneaaiiaeia?io (Eeyooi??i ? ?i., 1980). Ai?no
oinoao?a (caaaeueiiai, iai?aai?/iiai, ca’ycaiiai a
2,3-aeeoinoiae?oea?ao ? ioeeaioeaeiiai) aecia/aee iaoa?iaioiei iaoiaeii
ca Aeiia?aaeiaith ?.E. ? ?i. (1990). Aeey aecia/aiiy ai?noo AOO, AAeO
? AIO aeei?enoiaoaaee iaoiae ?acae?eaiiy a oiieiio oa?? nee?eaaaeth ia
ieanoeieao ‘‘Silufol’’ UV-254 (*ao?y). Aeey ioe?iee o/ano? ATO, AAeO,
AMO a iaoaaie?/i?e ?aaoeyoe?? ?ic?aoiaaii ni?aa?aeiioaiiy AOO/AAeO,
AOO/AIO oa aaeai?eaoiee aia?aaoe/iee ca?yae (Co?ae??, 1984).

Ai?no aetheice aecia/aee oioiiao?e/ii ca Anao?ai? A. N. (1957). ??aaiue
i??oaaoo oa eaeoaoo aecia/aee oa?iaioaoeaii ca I?ioi?iaith I. ?. (1982).
Ai?no a?eueieo IAAe-ia? a a?eo?ioeeoao ?ic?aoiaoaaee ia iniia? cia/aiue
eiioeaio?aoe?e noano?ao?a eaeoaoaeaa?ae?iaaiacii? nenoaie (*aaiaaoeue ?
?i., 1974).

Aeoeai?noue aetheici-6-oinoaoace a e?ia? aecia/aee ca I?aoiae/ai A. I.
(1968). ??aaiue iai?aai?/iiai oinoi?o ioe?ithaaee ca O?nea-Noaaa?io
(1925). Aeoeai?noue eaeoaoaeaa?ae?iaaiace a e?ia? aecia/aee ye
?aeiiaiaeiaaii a Biochemica Information (1975), a
aetheici-6-oinoaoaeaa?ae?iaaiace – ca Bergmeyer H. U.(1963).

No?oeoo?i? iiaeeo?eaoe?? a?ee?a e?ia? ioe?ithaaee ca ci?iith a?aenioea
ae?eiaaieo a?ee?a aaiie?caoo a?eo?ioeeo?a oa ieacie ca?aeii
?aeiiaiaeaoe?e Aeaieeiai? E. A. ? ?i. (1991). Nooi?iue ai?aeoaaiiy
a?ee?a e?ia? ioe?ithaaee ca Iooe?iith I. A. (1979). E?euee?noue
iaca’ycaieo noeueoa?ae?eeueieo a?oi e?ia? aecia/aee ca Oieiia?aei A. O.
(1981). Aaoa?iaaii?noue aaiiaeia?io ioe?ithaaee oeyoii eiai
aeaeo?ioi?aco a IAAA (Nooiieeiia ? ?i., 1972).

E?euee?noue a?eeo aecia/aee ii Lowry O. H. at al. (1951). Iaea?aeai?
?acoeueoaoe ia?iaeaii noaoenoe/ii c aeei?enoaiiyi iaoiae?a aa??aoe?eii?
noaoenoeee oa ei?aeyoe?eiiai aiae?co (Eae?i, 1990).

?ACOEUeOAOE AeINE?AeAEAIUe ? ?O IAAIAI?AIIss

3. Neeaae ? ooieoe?iiaeueia aeoeai?noue e?ia? ?ea

a ia??iae ceiiaiai aieiaeoaaiiy

A oiiaao ceiiaiai aieiaeoaaiiy a i?aai?ci? ?ea ?icaeaa?oueny
oaiia?aoo?iee oa ?ano?niaeao?oeeoiee no?an (Ni?aa/ia, 1982).
A?aeiia?aeaeth ia iueiai ? ca?eueoaiiy i?ioyaii cei?ae? a 1,5 ?ace
ai?noo a?eo?ioeeo?a oa aaee/eie aaiaoie?eoo. Caaaeueiee ai?no
aaiiaeia?io cieaeo?ony a 2 ?ace, ui ei?aeth? c aeaieie i?i i?yio
caeaaei?noue ai?noo aaiiaeia?io a ei?iia a?ae oaiia?aoo?e aiaee
(Na?ioi?i, 1979). Oea iiynith?oueny cieaeaiiyi a 4 ?ace aeoeaiino?
i?oioiiae??aeueii? d-AEN, yea e?i?oo? oaeaee?noue a?ineioaco aaia.
Caaaeii, aeai? ci?ie iia??oothoue a ia??iae cei?ae? caaacia/aiiy oeaiei
eeniai.

A e?ioe? aa?aciy ia ii/aoeo ea?oiy ii?yae c cieaeaiiyi a 4 ?ace ai?noo
ieneoi?ie aaiiaeia?io ??aaiue aeacieneoi?ie a?aeiinii ethoiai a 3 ?ace
c?inoa?. Ai?no iaoaaiiaeia?io i?e oeueiio ca?eueoo?oueny a 2 ?ace, ui
ni?aa?aeiineoueny c ??ceei cieaeaiiyi nii??aeiaiino? aaiiaeia?io aei
eenith. C aeaieie ci?iaie ei?aeth? oaeaee?noue aeenioe?aoe??
ieneaaiiaeia?io. Iaeneiaeueia oaeaee?noue aeenioe?aoe?? aeyaeaia a
aeiaoi?, eiee nii??aeiai?noue ? i?i?iaeueiith, a i?i?iaeueia – a
/a?ai?, eiee nii??aeiai?noue ? iaeneiaeueiith. Naciiia aeeiai?ea
aeenioe?aoe?? ieneaaiiaeia?io i?aeoaa?aeaeo? i?aeaeuaiiy
nii??aeiaiino? aaiiaeia?io aei eenith a ia??iae cei?ae?.

Ioaea, ne?ae a?aei?oeoe aca?iica’ycie i?ae i?aeaeuaiiyi nii??aeiaiino?
aaiiaeia?io aei eenith, cieaeaiiyi ??aiy iaooi?ie oa c?inoaiiyi
ai?noo eiai ieneoi?ie. Cia/ia cieaeaiiy ai?noo iene- oa i?aeaeuaiiy
e?eueeino? iaooi?ie aaiiaeia?io, a oaeiae ci?uaiiy e?eai? aeenioeaoe??
ieneaaiiaeia?io ai?aai a e?ioe? cei?ae? na?ae/eoue i?i ?icaaeainoaaiiy
nenoaie o?ainii?oo eenith a oeae ia??iae.

Ine?eueee a?io?i?/ia aaeaioaoe?y cae?enith?oueny ye oeyoii
iiaeeo?eaoe?? iae?iiieaeoe, oae ? ia ??ai? ?aaoeyoi?ieo oaeoi??a
ee?oeie (Oi/a/ea, Niia?i, 1988), iaie aeine?aeaeaii ?aaoeyoe?th
ooieoe?iiaeueii? aeoeaiino? aaiiaeia?io ca aeiiiiiaith AOO, yeee ?
iniiaiei iiaeoeyoi?ii nii??aeiaiino? aaiiaeia?io ?ea aei eenith (Bunn,
1971). I?ioyaii cei?ae? ??aaiue inoaiiueiai cieaeoaaany a 2 ?ace, ui
ni?anoaaey?oueny n a?aei?/aiei iaie oaeoii ca?eueoaiiy nii??aeiaiino?
aaiiaeia?io ei?iia aei eenith. Oea ocaiaeaeo?oueny c cieaeaiiyi a ?ea ca
iecueeeo oaiia?aoo? iai?io ?a/iaei, ?ooeeaino?, ciaioaiiyi niiaeeaaiiy
ieie eenith oa aeoeaaoe??th ae?eie?co.

Iaie aeyaeaii, ui c eenoiiaaea aei ethoiai a 1,3 ?ace ciaioo?oueny
??aaiue aetheice. I?e oeueiio aeoeai?noue ae?eie?co caaaeii c?inoa?,
e??i n?/iy, eiee a ?ea, ye a?aeiii (AEeaeaiei, 1992), aeoeao?oueny
aetheiiaiaaiac. Aiaeia?/ii ci?ith?oueny ni?aa?aeiioaiiy IAAe+/IAAeI,
yea ? ?aaoeyoi?iei oaeoi?ii ae?eie?co, aetheiiaiaaiaco oa aoaeaaiaeiiai
iai?io a oe?eiio. Oea ei?aeth? c aeoeaaoe??th a?ineioaoe/ieo i?ioean?a
a n?/i? oa ieenithaaeueii? caeaoiino? e?ia? a ae?oa?e iieiaei? cei?ae?.
Cieaeaiiy aeaiiai ni?aa?aeiioaiiy a na?aaeei? cei?ae? a?aei?/aii ia
o?eueee a e?ia?, aea e o i’ycao, ia/?ioe? oa iiceo ?ea (Eaeeie/aaa,
1974) ? nnaaoe?a (Aaeeeee ? ?i., 1982). Aeacai? caeiiii??iino?
i?aeoaa?aeaeothoueny oaeiae a?aeiia?aeieie ci?iaie aeoeaiinoae
eaeoaoaeaa?ae?iaaiace ? aetheici-6-oinoaoaeaa?ae?iaaiace oa aeeiai?eith
AOO, AAeO ? AIO a e?ia? ?ea. Noia aaeai?eiaeo ioeeaioeae?a oa
aaeai?eaoiee aia?aaoe/iee ca?yae i?ioyaii cei?ae? ciaioothoueny.
Cacia/aia ni?aa?aeiineoueny c a?aeiiei oaeoii aia?aicaaacia/aiiy
i?aai?cio ?ea a i?ioean? cei?ae? ca ?aooiie aiaeiaaiieo ?ano?n?a
(Ni?aa/ia, 1982).

Ia?aaoaeiae aia?aaoe/iiai iaoaaie?cio a?aeia?aaeathoueny ia
ooieoe?iioaaii? nenoaie o?ainii?oo eenith oeyoii aieeao ia
no?oeoo?ii-ooieoe?iiaeueiee noai aaiiaeia?io. Iaei??th c nooo?aeo
oa?aeoa?enoee inoaiiueiai ? e?iaoeea eiai eoaeii? aeaiaoo?aoe??.

Iaeno?ee?oei aaiiaeia?i ? a /a?ai? (ia??iae aeoeaiiai aeeaeaiiy).
Iaecia/i?oith eiai aeaiaoo?aoe?y ? a aeiaoi? oa ethoiio, ui ?
iieacieeii iecueei? caoeuaiino? iieaeoe a?ae aeaiaoo?oth/eo aaaio?a.

A oi?ioaaii? aeueo ??ai?a no?oeoo?e a?ee?a aaaeeeao ?ieue a?ae?a?athoue
ai?aei? a?oie (Aa?oaiiae/ ? ?i., 1966). I?ioyaii cei?ae? ai?aeiaai?noue
aaiiaeia?io i?aeaeuo?oueny a 2 ?ace. Ai?aeiaai?noue a?ee?a ieacie e?ia?
cieaeo?oueny a na?aaeei? cei?ae?, aea c?inoa? c ethoiai ii ea?oaiue.
A?aeiii (Eaeeie/aaa, 1974), ui c ae?oai? iieiaeie cei?ae? ye
aiaeiaaiia aeaea?aei aeeaeaiiy a ?ea aeei?enoiaothoueny naia a?eee, a
ca’yceo c /ei ai?aeoaaiiy aaiiaeia?io oa a?ee?a ieacie i?ioyaii
cei?ae? iiaeia aaaaeaoe caniaii ?o aaeaioeaiiai caoenoo a?ae
i?ioaie?co. Aeiaeaoeiaei iaoai?ciii oaeiai caoenoo ? ae?eoaaiiy
(Aeaieeiaa, 1991). Iaie aeyaeaii, ui a na?aaeei? cei?ae? ae?eoaaiiy
aaiiaeia?io cieaeo?oueny, a a aa?aci? c?inoa? iaeaea a 3 ?ace. A?eee
ieacie a /ania?e aeeiai?oe? ae?eothoueny iaaiaee. Aaaaaea?ii, ui
nooi?iue ae?eoaaiiy iiaea aooe oaeoi?ii, yeee c iaeiiai aieo ca
iani?eyoeeaeo oiia noaa?e?co? a?eeia? iieaeoee, a c ?ioiai – ?aaoeth?
?ioaineai?noue aoaeaaiaeiiai iai?io oeyoii aeeo/aiiy iiiinaoa?eae?a c
ieenithaaeueiiai ioeo. Ca?eueoaiiy nooiaiy ai?aeoaaiiy a?ee?a
noi?iaiaeaeo?oueny cieaeaiiyi a?aenioea ?o ae?eoaaiiy ? iaaiaee.
Eiia??ii, ui ca ?aooiie oeueiai iaeneiaeueii aoaeoeaii cae?enith?oueny
noaa?e?caoe?y aaiiaeia?io. Iaeii/anii c oeei iaaio ?ieue a caoeno?
aaiiaeia?io i?ae /an cei?ae? ?ea a?ae?a?athoue ? noeueoa?ae?eeuei?
a?oie. I?ioa ?o ?ieue, ii??aiyii c iienaieie iaoai?ciaie, ieae/a,
ine?eueee ai?no –SH a?oi i?ioyaii cei?ae? ci?ith?oueny iaei. Iacia/io
?ieue a aeaiiio i?ioean? a?ae?a?a? oaeiae ? iaoaaiiaeia?i?aaeoeoaca
a?eo?ioeeo?a, aeoeai?noue yei? ceiith cieaeo?oueny a 2 ?ace, ui ei?aeth?
c c?inoaiiyi ai?noo iaoaaiiaeia?io.

Aeai? aeene-aeaeo?ioi?aco aaiiaeia?io a IAAA aeacothoue ia oa, ui
aeyaeai? iaoai?cie caoenoo caaacia/othoue i?aeo?eiaiiy oe?e?niino? eiai
iieaeoe, ine?eueee e?euee?noue o?aeoe?e a?eea noaiiaeoue 3, ye ? a
ii?i?.

Oaeei /eiii, a ia??iae ceiiaiai aieiaeoaaiiy iieaeoea aaiiaeia?io oa
iaoaaie?ci a a?eo?ioeeoao ci?iththoueny a iai?yieo caaacia/aiiy
i?aeo?eiaiiy no?oeoo?ii? oe?e?niino? oa ooieoe?iiaeueii? aeoeaiino? aeey
caaacia/aiiy aeinoaoiuei? aaeaioiaaiino? aei aeno?aiaeueieo oiia
cei?ae?.

4. NEEAAe OA OOIEOe?iiaeueia AEOEAI?NOUe E?IA? ?EA CA ?IOIENEEAOe?E

4.1 Ii?oi-ooieoe?iiaeueia oa?aeoa?enoeea e?ia? ei?iia i?e ?ioieneeaoe?yo

Ca?eueoaiiy e?eueeino? a?eo?ioeeo?a oa aaiaoie?eoo ia? i?noea ca ae??
?ii?a naeioeth ie?aii oa ni?eueii c ai?aeii, ?ii?a ia?aaioeth, i?ae? oa
oaiieo, ui noi?iaiaeaeo?oueny ciaioaiiyi ai?noo a ieo aaiiaeia?io. Ca
ae?? ai?aeo ie?aii oa ni?eueii c ?iiaie iaai?th, ia?aaioeth, ?ii?a
oeeieo a?aei?/aii cai?ioith aeeiai?eo. I/aaeaeii, ui ca ae??
oieneeaio?a, iniaeeai ai?aeo, ?ii?a iaoae?a oa oaiieo ia? i?noea
ii?ooaiiy neioaco aaiiaeia?io. I?i oea na?ae/eoue ciaioaiiy ca ae??
ai?aeo, ?ii?a naeioeth oa oaiieo aeoeaiino? i?oioiiae??eii? d-AEN. A
?ioeo aeiaaeeao noai aaiiaeia?io aecia/a?oueny ni?aa?aeiioaiiyi eiai
e?aaiaeieo oi?i. Cieaeaiiy ??aiy ieneaaiiaeia?io ia niinoa??aaee ca
ae?? ?ii?a naeioeth ie?aii oa ni?eueii c ai?aeii, ia?aaioeth, a oaeiae
oaiieo. Cai?ioi?e aoaeo iaa i?noea o?eueee ca aieeao ia?aaioeth ?
ai?aeo. Ai?no iaoaaiiaeia?io a a?eueoino? aeiaaee?a ci?ith?oueny ia
iacia/ii (3-5%). I?e oeueiio i?ae ??aiyie iene- e aeacieneoi?i
a?aei?/aii cai?ioith caeaaei?noue. Iaea?aeai? aeai? na?ae/aoue i?i
?iaeea?aeoaeuei?noue ae?? aeine?aeaeoaaieo ?a/iaei, iaeiae, ni?eueiith ?
oaiaeaioe?y aei i?aeo?eiaiiy ooieoe?eii aeinoaoiueiai ??aiy iniiaieo
eiiiiiaio?a aaiiaeia?iiai? nenoaie.

Eiiiainaoi?ii-aaeaioeaieie ?aaeoe?yie ia oiene/io ae?th iiaeia aaaaeaoe
ca?eueoaiiy /enea a?eo?ioeeo?a, aeoeaaoe?th a?eo?iiiaoe/ii? aeoeaiino?
oeaiei, c?inoaiiy aeoeaiino? iaoaaiiaeia?ia?aeiiaethth/i?
IAAeH-caeaaeii? iaoaaiiaeia?i?aaeoeoace, iaeiiai c iniiaieo oa?iaio?a
aeieoaaiiy ieeneaiiy aaiiaiai cae?ca.

Aiae?c e?eaeo aeenioe?aoe?? ieneaaiiaeia?io na?ae/eoue i?i cieaeaiiy
nii??aeiaiino? aaiiaeia?io aei eenith ca ae?? ai?aeo ie?aii oa ni?eueii
c ?iiaie iaai?th, ?ii?a ia?aaioeth ? i?ae?, a oaeiae noi?o? ?ii?a
ia?aaioeth ? ai?aeo. A oiiaao ae?? ?ii?a naeioeth, naeioeth c ai?aeii,
oeeieo, oaiieo, a oaeiae i?e ci?i? eeneioiino? na?aaeiaeua e?ea?
aeenioe?aoe?? ci?uai? ae?ai. A aeiaaeeo ci?uaiiy e?eaeo ai?aai ca
iecueei? nii??aeiaiino? aeacieneaaiiaeia?io aei eenith a e?ia?
ca?eueoo?oueny ai?no iaca’ycaiiai iieaeoey?iiai eenith (?50 c?inoa?),
aiane?aeie /iai ?icaeaa?oueny ?? iaaeeeoeiaa ieneaaiaoe?y, ui a oiiaao
aeoeaaoe?? ae?eie?co iiaea i?eaanoe aei iaeiio?ieueiaaiiai
aeei?enoaiiy eenith ia ia ieeniaiiy aia?aaoe/ieo noano?ao?a, a ia
ooai?aiiy ia?ieneaeieo iio?aeieo e?i?ae?a (Aeaaeeie?ia, 1972).
Iaea?aeai? aeai? ei?aeththoue c oaeaee?noth aeenioe?aoe??
ieneaaiiaeia?io, yea, a aeiaaeeo c?inoaiiy ?50, ca?eueoo?oueny. C
?ioiai aieo, c?inoaiiy nii??aeiaiino? aaiiaeia?io aei eenith ca ae??
naeioeth ie?aii oa ni?eueii c ai?aeii, oeeieo, oaiieo, i?aeeeneaiiy oa
caeoaeaiiy aiaee ?icaeyaea?ii ye caoeni? ci?ie, ni?yiiaai? ia
iaeo?eiaiiy iaiao?aeii? e?eueeino? ooieoe?iiaeueii aeoeaiiai
aaiiaeia?io, ine?eueee iaaeeeoeiaa ieneaaiaoe?y, e??i IIE, iiaea oaeiae
ni?eyoe ieeniaiith aaiiaiai cae?ca.

Oaeei /eiii, ca?eueoaiiy nii??aeiaiino? aaiiaeia?io aei eenith iiaea
aooe oaeoi?ii i?aeo?eiaiiy ooieoe?iiaeueii iaiao?aeiiai ??aiy a?eueiiai
eenith (ae?y ?ii?a naeioeth ie?aii oa ni?eueii c ai?aeii, ?ii?a oeeieo,
caeeneaiiy ? caeoaeaiiy aiaee, aieea oaiieo) oa caniaii caiia?aaiiy
iaeiio?ieueiaaieo ia?aeenieo i?ioean?a.

Ni?anoaaeaiiy ci?uaiiy e?eaeo aeenioe?aoe?? ieneaaiiaeia?io oa ??aiy AOO
a a?eo?ioeeoao i?aeoaa?aeaeo? eiai ?ieue ye iniiaiiai iiaeoeyoi?a
ooieoe?iiaeueii? aeoeaiino? aaiiaeia?io ?ea. Ine?eueee a?eo?ioeeoa?iee
AOO ooai?th?oueny aeeeie?oe/ii (Aeoaea??a, 1979), oi ci?ia oaeaeeino?
ae?eie?co iiaea aieeaaoe ia eeniaaio?ainii?oi? iiaeeeaino? e?ia?.

4.2 Noai aoaeaaiaeiiai oa aia?aaoe/iiai iai?io a e?ia? ei?iia i?e
?ioieneeaoe?yo

Ocaaaeueithth/e caaaeueio aeeiai?eo ??aiy aetheice, i??oaaoo, eaeoaoo, a
oaeiae ieeniaieo oa a?aeiiaeaieo oi?i i?eioeiai?aeieo eioa?iaio?a,
iiaeia aeae?eeoe aeae?eueea iai?yie?a ci?i aoaeaaiaeiiai oa
aia?aaoe/iiai iaoaaie?cio a e?ia? ei?iia i?e ?ioieneeaoe?yo. Ca?eueoaiiy
??aiy i??oaaoo c iaeii/aniei cieaeaiiyi ai?noo eaeoaoo i?e ae?? ?ii?a
i?ae?, noi?o? ai?aeo c ?iiaie naeioeth, c?inoaiiy i?e oeueiio ai?noo
ieeniaieo oi?i i?eioeiai?aeieo eioa?iaio?a na?ae/eoue i?i i?eai?/aiiy
ae?eie?co oa iaeii/ania iineeaiiy ?ioeo oeyo?a ieeniaiiy aetheice,
o/anoue i??oaaoo a aetheici-aeai?iiaiio oeeee?, ?ieue yeiai iieyaa? a
i?aeo?eiaii? aiiainoaoe/iiai ??aiy ai?aeo oa aetheice. ??aaiue
inoaiiuei? ca aeacaieo ?ioieneeaoe?e a??ia?aeii c?inoa?.

Iaeii/ania ca?eueoaiiy ai?noo i??oaaoo oa eaeoaoo aeyaeaii i?e ae??
ai?aeo oa ?ii?a oeeieo. Oea na?ae/eoue ye i?i aeoeaaoe?th ae?eie?co, oae
? i?i aeei?enoaiiy i??oaaoo a iaoaaie?ci?. Iaeiie/aiiy iiei/ii?
eeneioe i?e ?ioieneeaoe?? ai?aeii ? iaiao?aeiei aeey i?aeo?eiaiiy
eeneioii-iniiaiiai aaeaino e?ia? i?e iaaaciaoe? ?? caeoaeaiiy.

Cieaeaiiy ??ai?a ye i??oaaoo, oae ? eaeoaoo i?e ae?? ?ii?a
naeioeth, oaiieo oa ia?aaioeth ie?aii ? ni?eueii c ai?aeii iiaea aooe
?acoeueoaoii ciaioaiiy ?ioaineaiino? ae?eie?co. Ca ae?? ?ii?a
naeioeth ? oaiieo cieaeo?oueny a?aeiioaiiy IAAe+/IAAeI, ui na?ae/eoue
i?i i?eai?/aiiy ieenithaaeueieo i?ioean?a, a oaeiae i?i ci?uaiiy
??aiiaaae eaeoaoaeaa?ae?iaaiacii? ?aaeoe?? aa?e eaeoaoo, ui ei?aeth? c
c?inoaiiyi ??aiy aetheice ? i?aeoaa?aeaeo? aeoieo i?i ciaioaiiy ??
ae?eie?oe/iiai ?icuaieaiiy.

Aiae?c aeoeaiino? eeth/iaeo oa?iaio?a aoaeaaiaeiiai iai?io – EAeA,
A-6-O-ace e A-6-O-Ae-ace a e?ia? ?ea aeicaiey? a?aei?oeoe aeae?eueea
iniiaieo iai?yie?a ci?i a nenoai? ae?eie?c – aetheiiaiaaiac –
iaioicioinoaoiee ooio (IOO):

– aeoeaaoe?th an?o oeyo?a a?aei?/aii ca ae?? ai?aeo ? caeoaeaiiy
aiaee. I?e ae?? ai?aeo i?aeo?eiaiiy aenieiai ??aiy aetheice ?
ca?eueoaiiy ai?noo ieeniaieo oi?i IAAe i?aeoaa?aeaeo? aeniiaie A?oa?iei
A. A. (1995) i?i iineeaiiy ?ioieneeaoe?? ?ea ai?aeii
aetheici-aeai?iiaiai oeeeeo. Iane?aeeii c?inoaiiy a?aeiioaiiy
IAAe(O)+/IAAe(O)I ? oaeiae aeoeaaoe?y IOO;

– aeoeaaoe?y ae?eie?co ? aetheiiaiaaiaco oa i?eai?/aiiy
iaioicioinoaoiiai ooioa aeyaeaii ca ni?eueii? ae?? ai?aeo c ?iiaie
iaai?th. C oeei iiaea aooe iia’ycaii cieaeaiiy ??aiy aetheice;

– aeoeaaoe?th ae?eie?co ? iaioicioinoaoiiai ooioa aeyaeaii ca ae??
noi?oae ai?aeo c ?iiaie naeioeth oa ia?aaioeth, a oaeiae ?ii?a i?ae? ?
oeeieo. Iaea?aeai? aeai? na?ae/aoue i?i iacia/io ?ieue aetheiiaiaaiaco
ca ni?eueii? ae?? ai?aeo oa ?ii?a naeioeth, a oaeiae ?ii?a i?ae?;

– i?eai?/aiiy ae?eie?co ? aeoeaaoe?th aeaio ?ioeo oeyo?a a?aei?/aii ca
ae?? ?ii?a ia?aaioeth ? caeeneaii? aiaee. Ca ae?? ?ii?a ia?aaioeth
aeaia iaoaaie?/ia ci?ia iiynith? a?aeiiee oaeo cieaeaiiy ??aiy aetheice
a e?ia? (Neciiaiei ? ?i., 1980);

– i?eai?/aiiy ae?eie?co ? iaioicioinoaoiiai oeyoo oa aeoeaaoe?y
aetheiiaiaaiaco i?iyaeythoueny o?eueee ca ae?? ?ii?a naeioeth;

– aeoeaaoe?th aeeeie?co oa i?eai?/aiiy ?ioeo i?ioean?a anoaiiaeaii
i?e ae?? oaiieo. Oiane?aeie oeueiai a e?ia? iaeiie/o?oueny eaeoao,
cieaeo?oueny a?aeiioaiiy a?eueieo IAAe-ia?, a oaeiae aeei?enoaiiy
aetheice.

Ci?ie oaeaeeino? ? ni?yiiaaiino? ?aaeoe?e aoaeaaiaeiiai iai?io
aieeaathoue ia ooieoe?iioaaiiy nenoaie ?aneioaco iae?ia?a?/ieo niieoe,
iniaeeai, aaeai?eiaeo ioeeaioeae?a.

Aeyaeaii cieaeaiiy ??aiy an?o aaeai?eao?a a e?ia? ca ae?? ?ii?a
naeioeth ? oaiieo, ui na?ae/eoue i?i i?eai?/aiiy i?ioean?a
aia?aicaaacia/aiiy oa, eiia??ii, ?icaeoie iaoieia?? a i?aai?ci? a
oe?eiio. I?aeaeuaiiy ??aiy AOO ? AIO i?e iaeii/aniiio cieaeaii?
eiioeaio?aoe?? AAeO ia? i?noea ca ae?? ?ii?a ia?aaioeth. I?e oeueiio,
ye ? ca ae?? ?ii?a naeioeth oa oaiieo, cieaeo?oueny ??aaiue caaaeueiiai
oa i?ia?aeueiiai oinoi?o. Oaiaeaioe?y aei ca?eueoaiiy ??aiy AOO ? AAeO
oa cieaeaiiy ai?noo AIO a?aei?/aia ca ae?? ai?aeo ?acii c ?iiaie
iaai?th. I?e oeueiio ciaioo?oueny ??aaiue an?o oinoao?a. Cieaeaiiy
ai?noo AOO c iaeii/aniei c?inoaiiyi ??ai?a AAeO oa AIO niinoa??aa?oueny
ca ae?? ?ii?a naeioeth ni?eueii c ai?aeii, a oaeiae ?ii?a i?ae? ?
oeeieo. Aeai? ci?ie na?ae/aoue i?i ?ioaineaia aeei?enoaiiy AOO aeey
i?aeo?eiaiiy aaeaioeaieo iaoai?ciia aeeoo?caaacia/aiiy. A??ia?aeia
ca?eueoaiiy ai?noo AAeO ? AIO i?e aeoeaaoe?? ae?eie?co oaeiae ?
na?ae/aiiyi ?ic’?aeiaiiy ieeniaiiy oa noano?aoiiai oinoi?eethaaiiy.
Oea ei?aeth? c aeaieie Aa?auaa?ia A. I. ? ?i. (1980) i?i aeoeaoth/o
?ieue AAeO oa, iniaeeai, AIO (??aaiue inoaiiueiai ca ?ioieneeaoe?e
aeoaea aenieee) ia oaeaee?noue ae?eie?co. Aeeiai?ea aaeai?eao?a oaeiae
ni?aa?aeiineoueny c ?aaoeyoi?iith ?ieeth AOO uiaei ooieoe?iiaeueii?
aeoeaiino? aaiiaeia?io.

Cieaeaiiy ??ai?a AOO oa AAeO i?e iaeii/aniiio i?aeaeuaii? ai?noo AIO
a?aei?/aii o?eueee ca ae?? ai?aeo. A?aoiaoth/e eiai niaoeeo?/io ae?th
ia e?ia (Emerson et al., 1975; Wlasow at al., 1990; A?oa?iei, 1995),
iiaeia ia?aaeaa/eoe AOO-acia ?icuaieaiiy AOO aeey i?aeo?eiaiiy
eeneioii-iniiaiiai aiiainoaco i?e aeoeaaoe?? ae?eie?co (Iaeueie/oe,
1989). Aiaeia?/? ci?ie iathoue i?noea oaeiae i?e caeeneaii? aiaeiiai
na?aaeiaeua. I?e caeoaeaii? aiaee aeoeaaoe?y ae?eie?co caaacia/o? ye
iineeaia ooai?aiiy eaeoaoo, iaiao?aeiiai aeey aeo?eioaaiiy
aioo??oiueia?eo?ioeeoa?iiai ?I, oae ? iineeth? ?aneioac AOO oeei
oeyoii.

Oaeei /eiii, ca ae?? noi?o? ?ii?a naeioeth oa ai?aeo, ai?aeo ie?aii,
?ii?a i?ae?, oeeieo, i?e i?aeeeneaii? aiaee ia? i?noea aeeaiea
?icuaieaiiy aaeai?eao?a, ui i?eaiaeeoue ia o?eueee aei iia??oaiiy
aia?aicaaacia/aiiy, a oaeiae aieeaa? ia ooieoe?iioaaiiy
aaiiaeia?io. Aeai? ci?ie iiaeia ?icaeyaeaoe ye iaoieia?th. I?eai?/aiiy
aeoeaiino? aaeai?eaoii? nenoaie niinoa??aa?oueny ca ae?? ?ii?a
naeioeth ? oaiieo. O aeiaaeeao ae?? noi?oae ai?aeo c ?iiaie ia?aaioeth
oa iaai?th, a oaeiae i?e caeoaeaii? aiaee eiia??iei ? aioaaii?ci
oieneeaio?a, ai ??aaiue AOO nooo?ai ia ci?ith?oueny. ?iie ia?aaioeth
aeoeaothoue i?ioeane aia?aiooai?aiiy a a?eo?ioeeoao.

4.3 No?oeoo?i? iiaeeo?eaoe?? a?ee?a e?ia? ei?iia a aeno?aiaeueieo oiiaao

Ia iniia? aiae?co iaea?aeaieo aeaieo iiaeia cai?iiiioaaoe
ooieoe?iioaaiiy aeae?eueeio iaoai?ci?a caoenoo
no?oeoo?ii-ooieoe?iiaeueii? oe?e?niino? a?ee?a e?ia? ca ae??
no?an-oaeoi??a. I?e oeueiio ??aaiue iiaeeo?eoth/i? ae?? oaeoi?o oa oei
aaeaioaoe?? caeaaeeoue a?ae o?ceei-o?i?/ieo aeanoeainoae oieneeaioo oa
aeeaeie eiai a?ieia?/ii? ae??. Ca oeei i?eioeeiii aeine?aeaeai? oaeoi?e
iiaeia ?iciiae?eeoe ia 3 a?oie:

1. Ca ae?? ?ii?a naeioeth iniiaia ?ieue a caoeno? a?ee?a iaeaaeeoue
SH-a?oiai, ye? ca,ycothoue ?o oa cieaeothoue aeinooi aei
ooieoe?iiaeueii aeoeaieo oeaio??a. Aiaeia?/i? ci?ie aeeeeeathoueny ca
ae?? ai?aeo, ?ii?a iaai?th oa oaiieo. I?e ae?? ai?aeo
noeueoa?ae?eeuei? a?oie iiaeooue aeei?enoiaoaaoeny ye aeiii?e i?ioii?a
aeey eiai ia?aaiaeo a iaio oiene/io ?ii?ciaaio oi?io.

2. Ca ae?? ?ii?a i?ae?, oeeieo oa caeeneaiiy aiaee ooieoe?iio?
iaoai?ci, yeee iieyaa? o eiioi?iaoe?eieo ia?aaoaeiaao a?eeiaeo
iieaeoe. Aiie ae?aaeathoueny a ca?eueoaii? ai?noo noeueoa?ae?eeueieo
a?oi c iaeii/aniei cieaeaiiyi ai?aeoaaiiy oiane?aeie ao?aoe
eaaeia?ae?ie?ciaaieo a?oi ania?aa?io oa ?ioaineo?eaoe?? i?ioean?a
ae?eoaaiiy ai?iieeneioieo caeeoe?a. A oeeo aeiaaeeao iniiaiee caoeniee
yoaeo caaacia/o?oueny ca ?aooiie ae?eoaaiiy.

?iie ia?aaioeth aeoeaothoue ?icaeoie an?o o?ueio cacia/aieo i?ioean?a.

Ooieoe?iiaeueiith oa?aeoa?enoeeith iiaeeo?eaoe?? iieaeoe aaiiaeia?io ca
ae?? oieneeaio?a ? e?iaoeea eiai eoaeii? aeaiaoo?aoe??. O nnaaoe?a
i?aeaeuaiiy ai?noo eoaeii?acenoaioiiai aaiiaeia?io aeyaeaii i?e
aiai?/i?e a?iien??, aeeeeeaii? ?ioieneeaoe??th naeioeai oa niieoeaie
?ooo? (Naiai/aaa, 1972). Eiia??ii, ui caeaoi?noue aei aeaiaoo?aoe??
aaiiaeia?io caeaaeeoue naia a?ae eiai no?oeoo?ieo ia?aaoaeia. Ai?ae
cieaeo? no?ee?noue aaiiaeia?io aei ae?? 0,1 i eoao. Eiai noi?o c
?iiaie iaai?th i?aeaeuothoue oeth caeaoi?noue ia a?aei?io a?ae
aeiaaee?a ae?? ?ii?a naeioeth ie?aii oa a noi?o? c ai?aeii, a oaeiae
?ii?a i?ae? ? oeeieo. Aieea ?ii?a ia?aaioeth ie?aii ? a noi?o? c
ai?aeii, oaiieo, i?aeeeneaiiy oa caeoaeaiiy na?aaeiaeua i?eaiaeeoue aei
ca?eueoaiiy no?eeino? aaiiaeia?io aei ae?? eoao. I/aaeaeii, ui
no?ee?noue aaiiaeia?io aei iacai?ioiueiai ii?ooaiiy iaoeaii? no?oeoo?e
eoaii aecia/a? nooi?iue iiaeeo?eiaaiino? eiai iieaeoe. Iaeiaioith
no?ee?noth aaiiaeia?i ?ea aieiae?? ca ae?? ?ii?a aaaeeeo iaoae?a ?
ai?aeo, ui iiaea aooe iia’ycaii c eiaaeaioiith iiaeeo?eaoe??th iieaeoe
a?eeia ca ?aooiie ai?aeoaaiiy, ooai?aiiyi iaei?ic/eiieo
iaoaeii?ioa?iiaeo eiiieaen?a oa caeoaeaiiyi aioo??oiueia?eo?ioeeoa?iiai
na?aaeiaeua. Inoaii? ci?ith? ca?yae iieaeoee a?eeo oa eiai
ooieoe?iiaeueio aeoeai?noue. I?aeaeuaiiy no?eeino? aaiiaeia?io aei eoao
na?ae/eoue i?i caeaaei?noue aeaiaoo?aoe?? a?eeo a?ae oa?aeoa?o
oieneeaioa oa oi?ie eiai ciaoiaeaeaiiy a i?aai?ci?, a oaeiae i?i caoenia
cia/aiiy aeine?aeaeaieo no?oeoo?ieo iiaeeoeeaoe?e. Na?ae/aiiyi
inoaiiueiai ? oa, ui, iacaaaeath/e ia aeinoaoiuei oe?ieee niaeo?
oieneeaio?a, aeaeo?ioi?aoe/ieie aeine?aeaeaiiyie a IAAA iaie ia
aeyaeaii ci?ie o?aeoe?eiiai neeaaeo aaiiaeia?i?a ?ea ca ?ioieneeaoe??
?o i?aai?cio. Iiaeia cacia/eoe, ui aaiiaeia?i ?ea aieiae?? cia/iith
no?oeoo?iith oa ooieoe?iiaeueiith no?ee?noth aei ae?? aeno?aiaeueieo
oaeoi??a aiaeiiai na?aaeiaeua.

5. Aca?iica’ycie no?oeoo?ieo iiaeeo?eaoe?e oa ooieoe?iiaeueii?
aeoeaiino? aaiiaeia?iiai? nenoaie ei?iia

A aeine?aeaeaieo aa??aioao ?ioieneeaoe?e ia aeyaeaii iaeiioeiii?
?aaeoe?? iinoo?aineyoe?eieo ci?i a?ee?a e?ia? oa ooieoe?iiaeueii?
aeoeaiino? aaiiaeia?io. I?e oeueiio a?aei?/a?ii, ui a aeine?aeieo oa
eiio?ieueieo aa??aioao e?euee?noue iiceoeaieo oa iaaaoeaieo ei?aeyoe?e
a?aei?iia ( +0,5

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